Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan

MIKHAIL A. AKHMETIEV

Paleocene and Eocene floras of Russia and adjacent regions are reviewed with an interpretation of climatic conditions under which they developed. Floristic and climatic changes in western and central regions of Russia in the Paleocene and in the first part of the Eocene were caused by the dynamics and rearrangement of the systems of marine seaways: a longi- tudinal seaway, which connected the Arctic basin with the marginal seas of Northern Peri-Tethys (Turanian, South Rus- sian and others), and a latitudinal seaway, which connected the marginal seas of Northern Peri-Tethys with the Atlantic Ocean. As these systems were progressively reduced, the climate in the middle latitudes changed from paratropical (like in West and Central Europe) to a subtropical monsoon climate with wet summers (Late Ypresian to Lutetian), and later to a climate with wet winters (Late Lutetian to the first part of the Priabonian). Floristic changes reflect these climatic trends. In the Paleogene, cold currents constantly influenced the climate of regions of the northwestern Pacific and facili- tated development of a warm-temperate mesophilic flora. A warmer episode took place in the Early Eocene. At this time thermophilic plants (Sabal, Myrtaceae and Lauraceae) reached Koryakia, North Western Kamchatka, probably as a re- sult of northward migration. Some subtropical plants existed near the Recent Polar circle. The subtropical Raichikha-type Flora lost temperate elements and, by ecological and climatic types, it is more similar to the Recent flora of South-West and South China. • Key words: flora, palaeoclimate, Paleocene, Eocene, Russia.

AKHMETIEV, M.A. 2010. Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan. Bulletin of Geosciences 85(1), 77–94 (7 figures). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received May 6, 2009; accepted in revised form October 5, 2009; published online January 8, 2010; issued March 22, 2010.

Mikhail A. Akhmetiev, Geological Institute of the Russian Academy of Sciences, Pyzhevsky by-street 7, 119017 Moscow, Russia; [email protected]

The quality of information about the Paleocene and Eo- Pacific. Onoclea, Osmunda, Ginkgo, Taxodiaceae, Cup- cene floras in Russia is very variable. Little is known ressaceae, Trochodendroides, Cornaceae, Hamamelida- about the floras from the northern and central parts of the ceae, Nyssaceae (Akhmetiev et al. 2002) and some other East European and Siberian platforms. Most localities are taxa are the most typical members of this flora. Trochoden- distributed along the coast of the Northern Peri-Tethys droides is a key indicator taxon for these floras (Fig. 1 – seas (South Russian, Turanian, West Siberian) and along Tr). To date, at least 70 species of the Tsagayan flora have the West Pacific periphery. Plant macrofossils are usually been recognized. Contemporary floras of the Tsagayan preserved in lacustrine and alluvial sediments of coastal ecotype are known from areas adjacent to the Amur River plains or, more rarely, in intermontane coal-bearing sedi- Area: southern Primorye, Sakhalin, Sikhote-Alin volcanic mentary basins as well as around volcanic structures. belts (Malo-Mikhailovka and Takhobe localities), Kam- Figs 1–4 show the main localities yielding plant macro- chatka and Koryakia (Fig. 1, locs 7–10). These floras are fossils. Seas, straits and key palaeogeographic areas are discussed in the final section of this paper. named in Fig. 2. At high latitudes, the warm temperate humid flora mi- grated westward along the northern boundary of the West Siberian plate and in the Danian reached the northern and Danian floras (Fig. 1) Middle Urals (Lozva flora, Fig. 1, loc. 1). The same type of the Danian flora occupied the Arctic region: Spitzbergen, At the K/T boundary, a transition to the warm temperate Greenland, North Canada, Alaska and northern Siberia – Tsagayan flora was first studied from the Tsagayan Series many localities at the Lena River Mouth area and of the Zeya-Bureya sedimentary basin (Fig. 1, locs 5, 6) Kharaulakh ridge (Bykov arm of the Lena River, Kengdey that developed on the continental boundary of the Western and Kungi grabens) (Budantsev 1983). Tsagayan plants

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Figure 1–4. Palaeogeographic maps and main localities for Paleocene and Eocene floras (within the boundaries of the former USSR). Legend (see Fig. 1): 1 – land, 2 – sea, 3 – locality and its number on maps, 4 – areas of bauxitization, 5 – direction of sea currents, 6 – plant indicators: Tr – Trochodendroides, U – Ushia, S – Sabal, N – Nypa, P – Palibinia. All geographical and paleogeographical elements are shown in Fig. 2. Figure 1. Early Danian. Localities: 1 – Lozva (western border of Western Siberian Plate) (Tr), 2 – Ulken-Kalkan (Ili River Basin, Kazakhstan) (Tr), 3 – Taizhusgen (Zaissan Lake basin) (Tr), 4 – Kara Biryuk (Zaissan Lake basin) (Tr), 5 – Raichikha coal field and Belaya Mount (Tsagayan flora – type locality) (Tr), 6 – Arkhara-Bouguchan coal field (Tr) (locality 5 and 6 – Zeya-Bureya sedimentary basin, Amur River Area), 7 – Malo-Mikhailovka (Lower Amur Area) (Tr), 8 – Kama River Area (Western Sakhalin) (Tr), 9 – Sobolevka (= Takhobe) (Eastern Sikhote-Alin) (Tr), 10 – Zerkalnaya (= Tadushi) (Eastern Sikhote-Alin) (Tr), 11 – Rarytkin Ridge (Northern Koryakia) (Tr), 12 – Beringovsky (Beringian sea, Anadyr Bay) (Tchukotskaya Formation) (Tr), Svb – Sikhote-Alin volcanic belt. migrated westward along the mid-latitude tracts from the the Late Paleocene. For each stage, only five to seven new Amur River system and reached Mongolia, Dzhungarian taxa (up to 10% of total) appeared, and approximately the Alatau, Ili River area (Ulken Kalkan), Semipalatinsk dis- same number of taxa disappeared. trict on the Irtysh River area and the Zaissan Lake basin There is the typical Danian Flora in the central part of (South Eastern Kazakhstan) (Fig. 1, locs 2–4). In the the Rarytkin Ridge (North Koryakia, Fig. 1, loc. 11) and Danian the Kara-Biryuk flora (Fig. 1, loc. 4) in the Zaissan a similar flora from Sakhalin (Kama River beds, Fig. 1, Lake basin includes typical Tsagayan plants: Taxites loc. 8) (Kodrul 1999). olrikii, Trochodendroides arctica, Nyssa bureica, Platanus raynoldsii, Porosia and others. In the Zeya-Bureya sedimentary basin (area of locali- Later Paleocene and Eocene floras ties 5 and 6 in Fig. 1), the Tsagayan flora went through of Northern Central Eurasia three stages of evolution from Middle Tsagayan to Kivda time (early Danian–?Selandian) (Kivda Flora see Fig. 1, Two main marine systems influenced the development of loc. 5). The youngest flora of the Tsagayan ecotypes is the Paleogene flora and vegetation in European Russia and characterized by various new members initially Myri- Siberia. 1) A meridional system that connected the Tethys caceae and Betulaceae as well as Ulmaceae. It should be with the Arctic Basin via the West Siberian epicontinental noted that no principal changes in composition of the sea and the Turgai strait. 2) A latitudinal system that con- Tsagayan flora took place in the Danian and possibly, in nected the East Peritethys with the Atlantic Ocean through

78 Mikhail A. Akhmetiev • Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan

Figure 2. Late Thanetian–Early Ypresian (including the PETM Episode). Localities: 1 – Kanev (Dnieper River Area, Southern Kiev), 2 – Kamyshin (Volga River Area (U, ?N), 3 – Nikulino-Privolsk (Volga River Area, northern of Saratov) (U), 4 – Akshuat – Tashly (northern of Saratov) (U), 5 – Smolino (South Ural, near Tchelyabinsk), 6 – Romankol (Or River Area, Northern Kazakhstan) (U), 7 – Tykbutak (Mugodzary Mounntain, Northern Kazakhstan) (U), 8 – Kiin-Kerish (Zaissan Lake Basin, brown clays “flora”), 9 – Kengday and Kunga troughs (Lower Lena River Area) (Tr), 10 – Bykovo channel (Lower Lena River Area) (Tr), 11 – Tas-Takh Lake (northern part of Siberian platform) (Tr), 12 – Beringovsky (Beringian Sea, Anadyr Bay) (Sandstones Member with ?Sabal), 13 – Tchemurnaut (Western Kamchatka) (Tr), 14 – Rebro Cape (Western Kamchatka) (Tr, S), 15 – Raichikha (Raichikhinsk coal field, Zeya – Bureya sedimentary basin), 16 – Rechnoi peninsula (northern of Vladivostok), Lower coal-bearing Member (Tr), 17 – Maikhe coal-bearing field (eastern of Vladivostok) (Tr). Areas of bauxitization: A – Arkalyk, B – Yenisei-Angara Area, C – Lower Yenisei River Area. Uk – Ukraine, Kz – Kazakhstan, K – Koryakia, Km – Kamchatka, SPV – South Primorie, Vladivostok, A – Amur River basin, Tm – Turkmenia. Bs – Barents sea, NZs – “Novaya Zemlya” strait, SZs –“Severnaya Zemlya” strait, Khs – Khatanga strait, WSs – West Siberian sea, Ks – Kara sea, Pb – Pechora bay, Ts – Turgayan strait, EEs – Eastern European sea, Tus – Turanian sea. Ps – Pripyat strait, Svb – Sikhote-Alin volcanic belt. the Dnieper-Donetsk Sea and western (Pripyat and Polish) Dewalquea, Macclintockia, Dryophyllum and Lauraceae. straits (Akhmetiev & Beniamovski 2006). This assemblage was found in several localities from the Volga basin in the Ulyanovsk and Saratov districts (Fig. 2, locs 2–4). The Kamyshin Flora contains Ushia in different Paleocene and Early Eocene evergreen morphotypes, which are described as species: Ushia kamy- thermophilic flora west of the Urals schinensis (Goepp.) Kolakovsky, U. alnifolia Makulbe- (Fig. 2, localities 1–4) and Kazakhstan kov, U. janishevskii Makulbekov, etc.) as well as Laura- (Fig. 2, localities 6–8) ceae (Daphnogene lanceolata Ung., Litsea sp., Ocotea sp., Persea paleomorpha Sap., P. enormis (Krasnov) Since the Danian, a Tethys Gelinden ecotype thermophilic Makulbekov, Dewalquea gelindensis Sap. & Mar., Oxy- flora, with mostly evergreen aspects, developed to the west carpia bifaria Tautsch., Ficus murczisonii Makulbekov, of the Ural mountain range under a humid “paratropical” Viburnum volgensie Krasnov, and others (Krasnov 1910, climate. Makulbekov 1977, Mai 1995). In the Late Paleocene the During the Selandian and the Thanetian the core of the Ushia-type flora spread to North Kazakhstan. The paratropical flora was formed by the Ushia assemblage Romankol flora (Fig. 2, loc. 6) from the Paleocene-Eocene (Fig. 2 – localities with U) with Chamaecyparis, Ushia, transition (corresponding with climatic optimum – PETM)

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Figure 3. Late Ypresian–Early Lutetian. Localities: 1 – Mogilno (western of Kiev) (S), 2 – Kremyanka (western of Kiev) (S), 3 – Zvenigorodka (Yekaterinopolye, Dniper River Area, south-eastern of Kiev), (Bouchak Formation) (S), 4 – Adzhamka (Dnieper River Area, south-eastern of Kiev) (Bouchak Formation) (S), 5 – Karakol (Mugodzhary Mountain, Northern Kazakhstan) (U), 6 – Chakelmes (Zaissan Lake Basin), 7 – Kershi (Zaissan Lake Basin), 8 – Karasor (North-Western Kazakhstan, Irtysh River Area) (?S), 9 – Northern Koryakia (Circus Mount) (Tr), 10 – Kangpil Cape (Penzhinsky Bay) (Tr), 11 – Snatol (Palana) (Western Kamchatka) (Tr), 12 – Due (Western Sakhalin) (Conglomerate and Lower Due formations) (Tr), 13 – Snezhinka (Lower Snezhinka Subformation) (Western Sakhalin) (Tr), 14 – Krasnoyarka River (Naibuti Formation) (South-Eastern Sakhalin) (Tr), 15 – Artyem Coal field (north-western of Vladivostok) (Uglovoye Formation) (Tr). included Ushia and various evergreen plants belonging to phyllum, Apollonias, Daphnogene, Phoebe), Ficus sp., Moraceae (Artocarpus, Ficus), Proteaceae (Dryandra), Ushia sp. (several morphotypes), Myrica banksiaefolia Magnoliaceae, Lauraceae (6 genera), Euphorbiaceae, Ung., Leucothoë protogaea (Ung.) Schimp., Bumelia ob- Elaeocarpaceae, Sterculiaceae, Combretaceae, Myrtaceae, longifolia Ett., Mimosites sp., Callistemophyllum mela- Theaceae, Melastomataceae, Araliaceae, Sapotaceae leucaeforme Ett., Sapindus sp., Aralia venulosa Sap. & (Sideroxylon), Symplocaceae, Apocynaceae etc. (Makul- Mar., Celastrophyllum sp., (?)Dryandra karakulensis Ba- bekov 1977, Baikovskaya 1984, Akhmetiev 2007). ranova, Poaceae gen., Dewalquea gelindensis Sap. & Mar. The Tykbutak locality (Fig. 2, loc. 7, Mugodzary and others. Mountain, Northern Kazakhstan) was studied by N.M. Locality 8 (Fig. 2) is situated on the northern Zaissan Makulbekov (1977). The fossils were collected from the Lake, on the western part of the Kiin-Kerish hill. Iljinskaya Tykbutak Formation. This formation consists of alternate (1963), Romanova (1970) and Akhmetiev (1985) studied sands, sandstones (predominate), sandy clays, and a form “Brown clays flora” at the base of the North-Zaissan of chert (a fine grained porous siliceous sedimentary rock Group. The composition of this flora changed from base to with small proportions of sand) [the Russian term for the top of the sequence. Near the base aquatic or semi-aquatic latter is opoka or gaises which seems to have no exact Eng- plants predominated, such as Peltandra, Zingiberopsis, lish equivalent]. Most of the leaves have entire margins and Nelumbo, Salvinia, Anoectomeria brongniartii Sap., Nym- belong to subtropical or tropical genera and families: phaea polyrhiza Sap., Carex sp., Cyperacites sp., Poacites Quasisequoia couttsiae (Heer) Kunzmann, Glyptostrobus sp., Phragmites sp. The rarer Protoacerophyllum perfo- sp., Magnolia embica Makulbekov, different Lauraceae liatum Romanova is also found near the base. Some aquatic (dominants, belong to Persea, Litsea, Laurus, Lauro- plants are like those from the Early Eocene Raichikha flora

80 Mikhail A. Akhmetiev • Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan

Amur River basin (Salvinia, Peltandra, Zingiberopsis, Ne- Lithocarpus, Dryophyllum and large-leaved Quercus pre- lumbo, Nymphaea). Both floras are characterized by reduc- dominated and also Magnolia, Sterculia, Leguminophyl- tion in conifers. This is a typical oxbow-lake assemblage. lum and other woody elements. Such assemblages were To the top of the section aquatic or semi-aquatic plants are documented by Makulbekov (1972, 1977) at Karakol reduced and different narrow leaved angiosperms appear, (Cis-Irtysian) (about 50% Castanopsis) and Sarkol (Mu- some of them endemics: “Begonia” vachrameevii Roma- godzhary) (both Fig. 3, loc. 5). Floristic layers with Cas- nova, “Oxicarpia” microcarpa Romanova, Protophyllum tanopsis overlap with the Lower Member of the Tykbutak zaissanicum Romanova, Arthollia (Zaissania) spp. Among Formation with Ushia. narrow-leaved endemic species there are Dryophyllum curticellense (Wat.) Sap. & Mar., Lindera vassilenkoi Iljin- skaya, Lauraceae gen., Apocynophyllum iljinskajae Roma- Late Ypresian–Lutetian, Ukraine (Fig. 3, nova and others. localities 1–4), North-Eastern Kazakhstan In Western Ukraine the Kanev Formation (Fig. 2, (Fig. 3, locality 8), Zaissan Lake Basin loc. 1) included clays and marls with rich Early Ypresian (Fig. 3, localities 6, 7) microfossil assemblages (planktonic foraminifera, nanno- plankton, dinocysts) distributed along the Dnieper valley To the west in Ukraine a large-leaved flora similar to that in from South to North Kiev. A.N. Kryshtofovich described Central Eurasia was described from the coal-bearing Bu- several species: Posidonia parisiensis (Desm.) Frittel, Aru- chak Formation (Lutetian). Most localities are concentra- nidites roginei (Wat.) Frittel, Macclintockia kanei (Heer) ted south of Kiev, Volynian (Mogilno locality, Fig. 3, Sap., Hakea spathulata Schmal., two small leaves similar loc. 1), and Dnieper River areas [Kremyanka (Fig. 3, locs to Lauraceae genera and Andromeda sp. (Leucothoë sp.) 2–4) (N), Jablontsy localities and others] (Schmalhausen (see Baranov 1959, p. 71). 1883–1884, Kryshtofovich 1927, Pimenova 1937, Stani- Bauxite (another indicator of warm wet subtropical cli- slavsky 1950, Zhilin 1986). Sabal (Fig. 3 – S) is an impor- mate) occurs in the Early Eocene in different areas of tant consituent of these floras. The main components of Kazakhstan and Siberia (Fig. 2). These include the areas these floras were Lauraceae (Daphnogene, Cinnamomop- near the Turgai-Arkalyk district of North Kazakhstan, hyllum, Laurus, Persea), Fagaceae (Dryophyllum, Eotri- along the Siberian platform (right side of the Yenisey River gonobalanus, Lithocarpus, Castanopsis), evergreen plants valley) from south to east along the south part of the represented by Ficus, Palmae (Sabal and Nypa), Magnolia, Yenisey ridge (60° north latitude) up to 67° N. Some stud- Myrtophyllum, Myrsine, Pittosporum etc. Ferns (Lygo- ies show that the ore was found near the Yenisey River dium) and conifers (Doliostrobus, Quasisequoia) were mouth (70° N) (Bushinsky 1958). very rare. Early Lutetian clays with Spondylos can be follo- wed along the right side of the Dniper Valley from Kiev to Yekaterinopolye (Fig. 3, loc. 3). Several localities with Late Ypresian subtropical humid forests, Nypa burtinii occur in this zone together with the typical central Eurasia (Fig. 3, locality 5) mangrove fern Acrostichum. Two localities with Late Ypresian–Early Bartonian flo- By the Late Ypresian the central Eurasian paratropical ras were found near the Zaissan Lake (South-Eastern climate changed to a summer-wet monsoon-type of sub- Kazakhstan). The first of them is Chakelmes hill (Fig. 3, tropical climate. This change was probably connected loc. 6). Dominants of the Chakelmes flora are Taxo- with interrupted latitudinal communications between the diaceae, Dryophyllum curticellense (Wat.) Sap. & Mar., D. western and eastern North Peri-Tethys seas. A land bar- dewalquei Sap. & Mar., D. kryshtofovichii Iljinskaya, dif- rier was formed on the Pripyat strait (Akhmetiev & Benia- ferent Lauraceae (dominant, more then 80% specimens, movski 2006). By the Early Lutetian the inner marine seas determined by epidermal investigation) and Zingiberopsis began to shallow and showed signs of reduction of the sp. More rarely encountered are Sequoia sp., Taxodium longitudinal straits connecting the Arctic basin with the dubium (Sternb.) Heer, Glyptostrobus sp., Cryptomeria Tethys through theTurgai strait and West Siberian Sea. sp., Magnolia sp., Populus kryshtofovichii Iljinskaya, The Orsk-Ayat strait in the southern Ural disappeared. Lindera sp., Apocynophyllum sp. and others (Iljinskaya The West Siberian Sea also reduced, but southern transfer 1986, Akhmetiev 1993). of heat and moisture continued elsewhere. Seasonal redis- The second flora is from Kershi (a small hill in the tribution of precipitation took place. Early Paleogene pa- northern Zaissan Lake area) (Fig. 3, loc. 7). This is a typical ratropical forests changed to subtropical humid forests as sclerophyllous subtropical-type flora. E.V. Romanova and a result of the onset of a monsoon summer-wet climate. I.A. Iljinskaya recognized more than ten species: Liquid- These forests included Lauraceae (Persea, Litsea, Daph- ambar kazakhstanica Romanova, Ficus sp., Coccoloba sp., nogene) and Fagaceae, especially Castanopsis, and also Sterculia sp., Cassia sp., Eugenia kirshiensis Romanova,

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Idesia romanovii Iljinskaya, Ungnadia kryshtofovichii basins ceased. The warm current continued from Tethys to Iljinskaya and others (Iljinskaya 1986). the surviving large West Siberian marine gulf separated by The typical monsoon-type subtropical flora is found in a land barrier from the Arctic Basin. The climate changed Karasor (North Eastern Kazakhstan, Irtysh River area) and monsoon summer-wet climate was replaced by seaso- (Makulbekov 1972) (Fig. 3, loc. 8). The main components nal winter-wet climate, similar to that of the present region of this flora are Fagaceae, especially Castanopsis (more of the Mediterranean. than 50% C. kazachstanensis Makulbekov and C. pavlo- The floral lists herein are based on (i) the author’s own darensis Makulbekov), Quercus kornilovae Makulbekov, collection of the xerophytic flora from the Baky locality Q. irtyshensis Makulbekov, Quercus spp., Dryophyllum (Fig. 4, loc. 5), southern Ural, (ii) the author’s revisions of curticellense (Wat.) Sap. & Mar., Dryophyllum dewalquei other collections from Baky made by M.D. Usnadze-Dge- Sap. & Mar. More rarely found are “Podocarpus” eoce- buadze and N.M. Makulbekov and (iii) the author’s revi- nica, Lauraceae (Persea, Cinnamomum, Litsea), Myrica sions of collections from neighboring areas: Mugadzhary, spp., Leucothoe protogaea (Ung.) Schimp., Diospyros sp., Emba River area and Pavlodar district (Cisirtysh area), Sterculia kalkamanica Makulbekov, Leguminosae gen., which were studied by A.N. Kryshtofovich, V.I. Baranov, Eugenia haeringiana Ung., Lomatia macrophylla Makul- L.Y. Budantsev, I.V. Vassiliev and V.S. Kornilova during bekov, Echitonium sp., Palmophyllum sp. and others. the 1950–1970s. After the revision, the flora contains some conifers (“Sequoia”, “Podocarpus”) and angiosperms, such as Late Eocene, mixed evergreen deciduous, Macclintockia (= Majanthemophyllum), some Myricaceae, amber flora, Sambian peninsula, Baltic Region Fagaceae and Fabaceae. The typical Baky flora and other (Fig. 4, locality 22) floras of the same composition from the Or’ – Ilek water- shed (Romankol, upper floristic layers, Zharlykap, Fig. 4, The Late Eocene flora of the Sambian peninsula (Baltic loc. 6 with Sabal, Fagaceae gen. (evergreen), Leucothoë, Region, Kaliningrad district) is unusual. It is represented Palibina and others) and Or’ River, South Urals (Fig. 4, by equal amounts of evergreen and deciduous plants. It is loc. 7) with Cupressaceae and small sclerophyllous leaves preserved in pieces of amber as inclusions of flowers, fruits Angiospermae), Mugodzhary, Pavlodar district [Takyrsor and fragments of leaves. Fossil plant remains belong, besi- (Fig. 4, loc. 13), Zhaman Tuz (Fig. 4, loc. 12 – Makulbekov des angiosperms, to mosses, ferns, Pinus needles and rep- 1972] include Athrotaxis,?Doliostrobus, Cupressaceae roductive organs, Cupressaceae and rare Taxodiaceae. gen. indet, Laurophyllum, Myrica, Rhodomyrtophyllum, About two hundred species of angiosperms (especially fru- Quercus, Eotrigonobalanus, Leucothoë, Majanthemo- its and seeds) were described including evergreen Faga- phyllum, Leguminophyllum (different species), Echito- ceae, Lauraceae, Anacardiaceae, Hamamelidaceae, Aralia- nium etc. This list can be complemented by plants which ceae and others (Conwents 1886, 1890). This flora are not represented in our collection: Palibinia (small characterizes an ecotonal zone between the Boreal and Tet- sclerophyll narrow lobed leaves similar to those of hys realms. Comptonia or Dryandra), Sabal and some others (data from N.D. Vassilevskaya). Southern Uralian and Mugo- dzharian species of Palibinia match the Badkhys Palibinia Bartonian, sclerophyllous flora, Ukraine that Korovin (1932, 1934) described from the Bartonian to Kazakhstan (Fig. 4, localities 5–9, 12, 13), Turkestan regional stage. and Turkmenia (Fig. 4, localities 10, 11) The Akar-Cheshme flora (Western Badkhyz, Turk- menia) (Fig. 4, loc. 10) with Palibinia, was discovered by A xerophytic Bartonian flora is distributed in the latitudinal N.D. Vassilevskaya (1957). Fossil plants are concentrated zone from northern Ukraine through the Voronezh anti- in thin ash layers (Andesite-dacite tuffaceus Member) un- cline, southern Ural and Mugodzhary to eastern Kazak- derlying and overlapping marine clays with Bartonian hstan (Pavlodarian Irtysh River area). The localities often planktonic foraminifera and nannoplankton assemblages. contain Palibinia (Fig. 4 – P) as an important indicator ta- The Akar-Cheshme assemblage consists of different en- xon. Plant remains are preserved in red cavernous quartzite demic ferns and angiosperms conifers are absent. There sandstones and represent mostly chaotically twisted small were determined by N.D.Vassilevskaya: Blechnum akar- and narrow sclerophyllous leaves of angiosperms and coni- tscheschmens Vassilevskaya, Lastraea badchyzica Vas- fers. Empty holes in the rock resulted after plant tissues of silevskaya, Polypodites nepphrotiopsis Vassilevskaya, trunks and branches decayed after being covered by eolian Myrica dilodendrofolia Vassilevskaya (dominant), Carya sands and dust-like particles. The quartzite sandstones are typica Vassilevskaya, Banksia myricifolia Vassilevskaya, younger than the Late Lutetian just after the meridional Lauraceae (Ocotea, Laurus), Amygdalus turcmenensis Vas- marine communication between the Tethys and Arctic silevskaya, Leguminosae gen., Anacardites badchyzicus

82 Mikhail A. Akhmetiev • Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan

Figure 4. Late Lutetian–Bartonian. localities: 1 – Volyantschina (west of Kiev), 2 – Kiev (Lower Kiev Subformation) (N), 3 – Putivl (east of Kiev) (N), 4 – Akhaltsikhe (Georgia), 5 – Baky (Southern Ural) (S), 6 – Zharlykap (Or River Area, Southern Ural) (S, P), 7 – Or (Or River) (Southern Ural), 8 – Zhaman-Kaindy (Turgai District, Northern Kazakhstan) (P), 9 – Tortmolla (Turgai District, Northern Kazakhstan) (P), 10 – Akar-Cheshne (Western Badkhys, Turkmenia) (P), 11 – Er Oilan-Duz, Kushka district, Turkmenia) (P), 12 – Zhaman Tus (Irtysh River Area, North Western Kazakhstan), 13 – Takyrsor (Irtysh River Area, North Western Kazakhstan) (S), 14 – Tuzkabak (Zaissan Lake Basin), 15 – Kalmakpai (Zaissan Lake Basin) (Sargamyss Formation), 16 – Shestakovo (Penzhina Bay) (Tr), 17 – Podkagernaya (Western Kamchatka) (Irgirnin Formation) (Tr), 18 – Snezhinka (Western Sakhalin) (Upper Snezhinka Subformation), 19 – Sonye (Eastern Sikhote-Alin volcanic belt) (Tr), 20 – Rechnoi peninsula (northern of Vladivostok) (“Nadezhdinskaya” Formation) (Tr), 21 – Svetlovodnaya (Eastern Sikhote-Alin volcanic belt), 22 – Sambian peninsula, Prussian Forma- tion, amber flora, Late Eocene, Svb – Sikhote-Alin volcanic belt.

Vassilevskaya, Rhus sp., Zizyphus spp., Tetracera sp., Middle Eocene marine deposits with rich dinocyst assem- Melastomites sp., Diospyros sp. Palibinia is one of the blage (N.I. Zaporozhets, personal communication). Fossil dominant plants in this assemblage. plants are concentrated in fossil soil horizons on a red-or- The Er Oilan-Duz flora (Kushka district, Turkmenia, ange crust of weathering over marine deposits. The Fig. 4, loc. 11) has the same stratigraphical position. The Oligocene clays with Chalicotherium (V.Y. Reshetov, per- main components of this flora are Palibinia Korovin (sev- sonal communication) overlap and cross-cut the soil hori- eral species), Rhus turkomanica (Krysht.) Korovin and zon. Therefore, the Tortmolla flora with Quasisequoia Cinnamomum sp. Leaves of Rhus sp., Zizyphus and Dio- couttsiae, Palibinia, Rhus turcomanica, Leucothoë, spyros were also found. Both of the Bartonian Turkmenian Echitonium, etc. is probably of Middle or Late Eocene age. floras have small- and sclerophyllous leaves. The Tortmolla flora (only microphyllous) is a more The age of some Turgaian Paleogene floras, e.g. the xeromorphic type than the South Uralian and Cisirtyshian Tortmolla flora (Fig. 4, loc. 9) with Palibinia and Rhus floras [see Fig. 2, loc. 5 – Smolino (with Oxycarpia), loc. turcomanica and the Zhaman-Kaindy flora (Fig. 4, loc. 6 – Romankol, loc. 7 – Tykbutak; Fig. 3, loc. 5 – Karakol; 8), originally dated to Early Oligocene (Zhilin 1989), has Fig. 4, loc. 5 – Baky, loc. 6 – Zharlykap, loc.7–Or(All been revised. Zhilin (2005) changed his opinion and dated South Ural and Mugodzhary), loc. 8 – Zhaman-Kaindy, the Tortmolla flora to the Late Eocene. It is now possible loc. 10 – Tortmolla (both of Turgai, North Kazakhstan)]. to determine more precisely the stratigraphical position of Judging according to the distribution of recent vegetation the Tortmolla floristic layers. These layers overlap with analogues of the Bartonian xerophytic plant assemblages,

83 Bulletin of Geosciences • Vol. 85, 1, 2009 the precipitation was 200–600 mm per year, and the mean Kalmakpai area. It includes in addition Salvinia natanella, summer temperatures were 22–26 °C. Zonal types of veg- Craigia, Lindera vassilenkoi, and Ungnadia. The flora of etation north of the Peritethys coastal plane and southern the Tuskabak depression includes also Dryophyllum, Ni- coast of the West Siberian sea were sclerophyllous wood- tophyllites and Alnus. lands with different major components and rich in shrubs The Late Eocene flora of the Turanga Formation (like recent maquis) with narrow-leaved Ericaceae, (Kiin-Kerish hill) was studied by Iljinskaya (1963, 1986) Myricaceae, Fagaceae, Fabaceae, Myrtaceae, and sclero- and Akhmetiev (1985). Thin-bedded light yellow clays and phyllous conifers. thin sands (22–25 m) overlaid by variegated deposits with the Bartonian flora include three floristic layers of the Turanga flora (9, 12 and 20 m above the base of the Late Eocene freshening of sea surface waters Turanga Formation). Late Eocene swamp rhinoceros Zaissanamynodon borissovii and Kiinkerishella zaissanica At this time, the partly isolated West Siberian marine basin (Arctocionidae) were found 2 m above the base of this sec- opened to the Tethys. One of the main groups for biostra- tion (7 m lower than the first floristic layer). In the lower tigraphic dating is organic-walled phytoplankton indica- floristic layer Salvinia mildeana, Nelumbo, Zingiberites ting a mid to late Eocene age at the Bartonian-Priabonian borealis are dominant. “Hibiscus” neuburgae and Ulmus transition. At the beginning of the Late Eocene surface wa- are rarer. ters of the West-Siberian inner sea and Turanian sea on the The next floristic layer (3 m further above) is formed by North-Eastern Peri-Tethys margin were freshened after a clays. This riparian assemblage consists of Alnus, Ulmus, continental bridge formed at the southern Arctic interrup- Populus arnaudii,“Hibiscus” neuburgae, Zelkova zelko- ting the Arctic/Tethys connection. This process was repea- vifolia, rarely Lindera sp., Myrica sp., Leguminosae spp. ted several times. At the first stage layers with Pediastrum (leaves and fruits), Dryophyllum spp., Sterculia sp., Euca- were formed there, in the second stage layers with Hydro- lyptus sp. and Apocynophyllum sp. Several leaf beds were pteris indutus, and in the third stage layers with Azolla (me- formed by Ulmus sp. and “Hibiscus” neuburgae. gaspores and leaves) (Akhmetiev et al. 2004a, b; Akhme- The upper floristic horizon is formed by a lens of white tiev & Zaporozhets, personal observations). During the last clay just below the red soil horizon (first stage of originat- century also impressions of leaves of Azolla vera Krysht. ing crust of weathering). Fossil plants are sclerophyllous, were recovered in the lower parts of the Chegan and Tavda of middle size, some also narrow-leaved. The following formations (Bartonian-Priabonian transition) at many loca- elements were determined: Dryophyllum curticellense, D. lities (Aral sea cliffs, Turgai and Irtysh areas in western dewalquei, D. krystofovichii, Myrica acuminata, Myrica Pavlodar escarpments of cuestas, West Siberia, and in cf. lignitum, Taxodium dubium, Lindera vassilenkoi, South Siberian boreholes samples). Zelkova zelkovifolia, Ampelopsis schischkinii, Fraxinus, Astronium ninae, Craigia, etc. Middle and Late Eocene, mixed deciduous and evergreen floras, Zaissan Lake Basin Later Paleocene and Eocene floras (Fig. 4, localities 14, 15) in the Western Pacific Margin

The Middle and Late Eocene floras of the Zaissan Lake ba- The majority of the Western Pacific Paleogene plant locali- sin are characterized by a mixture of evergreen and decidu- ties are situated in intermontane coal bearing deposits in ous elements. The climate was more humid, but seasonal. depressions, some of them occupying vast areas (e.g., the Three localities of Bartonian age are known in this area: the Zeya-Bureya sedimentary basin, Amur River area) in ma- Kalmakpai (Fig. 4, loc. 15) and Kiin-Kerish hills (same rine coastal zones, and in sedimentary tuffaceous beds of area as locality 8 in Fig. 2 with a continuous section from the West Pacific volcanic belts. Late Paleocene to Oligocene) and the Tuskabak depression The main characteristics of the East Asian floras and area (Fig. 4, loc. 14) around the Zaissan Lake. Plant rema- climate are principally different from contemporaneous ins are concentrated in variegated clays between two red floras and climate in Central and Western Russian Eurasia weathering clayey crusts. The main components of the Mid at middle latitudes. The East Siberian and Far East to Late Eocene Kiin-Kerish flora (I.A. Iljinskaya and Akh- Paleogene floras belong to the Boreal realm. The compo- metiev, personal observations) are “Hibiscus” neuburgae, sition of East Asian floras was influenced by a cold cur- “Hibiscus” kalmakpaica, Populus kryshtofovichii, Faba- rent running along the eastern coast from Beringia ceae (leaves and fruits), rare Comptonia, small-leaved Ul- through Kamchatka, Sakhalin to Japan. The stability of mus, Liquidambar, Celtis, Zelkova, Magnolia, and Ailan- this current was maintained by the long existing Bering thus. The richest is the Sargamyss Formation flora in the Land Bridge.

84 Mikhail A. Akhmetiev • Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan

Early Eocene floras of the high latitude plants reaching 64%. Among them are Lygodium, Ocotea, Siberian Platform (Fig. 2, localities 9–11) Cinnamomophyllum, Corylopsis, Platycarya, Camellia, Byttneria, Dombeya, Myrtophyllum, Leguminosae (fruits The Early Eocene climate at the northern margin of the and leaves), and others. Leaves of Sabal were discovered Siberian platform was similar to the climate of the polar between the capes Rebro and Kengi (Fig. 2, loc. 14). A sea, where the surface-water temperature reached +20 °C partial specimen of Sabal (lacking the rachis) was found (Backman et al. 2006). It was frostless or with episodic by V. Volobuyeva in levels of the same age south of the short cold winters. This is confirmed by some localities mouth of the Anadyr River (North Koryakia) (Fig. 2, that are nowadays situated within the Arctic Circle, loc. 12). Unfortunately the specimen is now lost so the re- north of 70 N (grabens on the lower course of the Lena cord is treated as ?Sabal on the map. During the Middle River (Fig. 2, locs 9, 10) and Lake Tastakh (Fig. 2, Eocene cooling most thermophilic plants disappeared and loc. 11) in the Yana and Indigirka interfluve). These flo- the Irgirnin flora corresponds to a temperate mesophilic ras contain remains of gymnosperms (Ginkgo, Pinaceae, type, with numerous conifers (Pinus, Abies, Larix, Meta- Taxodium, Metasequoia, etc.), both deciduous broad- sequoia, Taxodium, Cryptomeria, Cunninghamia, Scia- leaved trees (Trochodendroides, Platanaceae, Ulma- dopitys, etc). The appearance of Fagus, Castanea and ceae, etc.) and thermophilic southern migrants, such as some other modern genera, analogous to the Irgirnin Myrtaceae and Liquidambar represented by leaves and flora, was recorded by the author from the Ust – Penzhina reproductive organs (Ozerov et al. 2006), leaves of Mag- Formation (Penzhinskaya Bay, South Koryakia), in the nolia, Palmae (pollen), Loranthaceae (pollen), and Ha- central part of the Rarytkin Ridge (North Koryakia) (for mamelidaceae (Kulkova 1973). location see Fig. 1, loc. 11 an area with both Danian and Early Eocene floras). T.M. Kodrul described a similar flora from Sakhalin (Kama River beds, Fig. 1, loc. 8) Early Paleogene floras of North East Russia (Kodrul 1999). (Fig. 2, localities 12–14, Fig. 3, locality 11), In Northern Vladivostok an Early Eocene flora was Sakhalin (Fig. 1, locality 8) and South Primorie found at the base of coal-bearing deposits on the Rechnoi (Fig. 2, localities 16, 17) peninsula (Uglovaya Formation, Taurichanka coal-field, Fig. 2, loc. 16). This is a typical Metasequoia-Trocho- The early Paleogene floras of northern Russia, like the dendroides-Platanaceae assemblage (Akhmetiev 1993). preceding Late Cretaceous flora, were broad-leaved and Most of the plants are concentrated in coal-bearing sand- existed under conditions of limited day-light during short stones and silts (middle part of Uglovaya Formation). In winter days within the Arctic Circle. L.Y. Budantsev addition to these dominants Magnolia sp., Pterocarya sp., (1983, 1986, 2006) described three stages of floral evolu- Zelkova kushiroensis Oishi & Huzioka, Nelumbo tion in the Early Paleogene of the western Kamchatka nipponica Endo, Hamamelis sp., Liquidambar sp., Rhus Peninsula (which he referred to as the Beringian Province sp. and Plafkeria sp. were identified. of the Boreal Realm), (i) Khulgun (Paleocene), (ii) Tche- In eastern Vladivostok in the Maikhe coal-bearing field murnaut (Fig. 2, loc. 13) (Early and Early Middle Eocene) (Fig. 2, loc. 17) some thermophillic plants [evergreen Fa- and (iii) Irgirnin (Late Eocene according to Budantsev, gaceae, Lauraceae and Leguminosae (Akhmetiev 1993)] Middle-Late Eocene transition according to Zaporozhets et were found at the base of the Uglovaya Formation. al. (2006) based on a dinoflagellate assemblage at the base of the Ratygin Formation which conformably overlies the plant-bearing Irgirnin Coal Formation). Eocene warm temperate floras The Khulgun-type flora (Palana locality, near Snatol) of the Russian Far East (Fig. 3, loc. 11) was rich with more than 70 species two thirds of which appear for the first time (Budantsev 2006). The Far East flora remained warm-temperate, meso- These include Osmundastrum, Dennstaedtia, Ginkgo, philic and deciduous till the Late Eocene and retained Magnolia, Lindera, Tetracentron, Betula, Corylus, Alnus, some Early Paleogene morpho-genera like Trochoden- Carpinus, Myrica, Juglans, Pterocarya, Acer, Hydrangea droides or Fagopsis. Only during global warming at the and others. The same type of Paleocene flora is known on Paleocene/Eocene boundary and in the Early Eocene did Sakhalin (Kama beds) containing Taxodiaceae, Ulmaceae, some evergreen immigrants invade into high latitudes. Craigia, Trochodendroides arctica and others (Kodrul At that time, palms (Sabal) reached Western Kamchatka 1999). The Tchemurnaut stage coincides with the Early and Koryakia (as far as to 64° N). The boundary between Eocene major global warming. The Tkapravajam Forma- the Boreal and Tethys realms was shifted northwards by tion flora (Budantsev 2006) includes at least 40% en- 8–10° in the Far East (Akhmetiev et al. 2002, Budantsev tire-margined leaves and a high diversity of thermophilic 2006).

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Eocene subtropical Raichikha flora the subtropical zone migrated at least 20° further north rel- (Fig. 2 locality 15, Figs 5–7) ative to its present position. The Early Eocene climate was subtropical monsoon (summer wet). This is confirmed for The Lower Paleogene deposits of the Zeya-Bureya sedi- the Raichikha Formation by active kaolinization. mentary basin include two types of floras: warm-temperate Tsagayan (Paleocene) (see section on Danian floras) and Raichikha (Eocene). The Early Eocene flora of the Rai- Late Ypresian and Lutetian floras chikha coal-field (50° N) has a distinctive composition, of the North-Western Pacific coast (Fig. 3, with typically subtropical elements and including various localities 9–11), Sakhalin (Fig. 3, localities evergreen plants (Figs 5–7). This flora was studied by T.N. 12–14), Primorie (Fig. 3, locality 15) Baikovskaya (1950). Later A.N. Krystofovich (1952) noti- ced the unusual composition of the flora. M.A. Akhmetiev The floras of the localities of the North-Western Pacific: (1969, manuscr.), V.V. Fedotov (1983, doctor thesis), and Circus Mount (North Koryakia, Fig. 1, loc. 9) (Akhmetiev A.M. Kamaeva (1990) described many new taxa. The Rai- 1993), Kangpil Cape (Penzinsky Bay, Fig. 3, loc. 10) (Akh- chikha and preceding Tsagayan floras have very few ele- metiev 1993) and Snatol River (Western Kamchatka, loc. ments in common (except for some ferns, Taxodium, Poro- 11) (Budantsev 1997, 2006) are similar and represent typi- sia and Platanaceae). The Tsagayan-type flora could not be cal Late Ypresian–Early Lutetian assemblages with ferns an ancestral assemblage to the Raichikha flora. The Rai- (Osmunda macrophylla Penhallow, Dennstaedtia ameri- chikha flora includes southern elements which might have cana Knowlt., Lygodium coloradense Knowlt.), Ginkgo migrated from the low latitudes during the Early Eocene kamtschatica Budants., Metasequoia occidentalis (Newb.) warming. Modern analogues of the Raichikha flora are wi- Chaney, Taxodium tinajorum Heer, Glyptostrobus europa- despread through South China and also Japan (30° N). The eus (Brongn.) Heer, Magnolia basicordata Fotjanova & O. Raichikha flora lacks palms, but it is characterized by di- Lavrenko, Trochodendroides arctica (Heer) Berry, Nyssi- verse ferns, various evergreen and deciduous angiosperms. dium arcticum (Heer) Iljinskaya, Cocculus ezoanum Tanai, This flora is composed of one member of Hepaticopsida, Nordenskioldia borealis Heer, Populus microdentata Bu- one Equisetum, 10 ferns, one conifer, more than 70 species dants., P. celastrophylla (Baikovskaya) Sycheva, Carpi- of dicots (of which 44.6% have entire margined leaves) and nus ex gr. C. japonica Blume, Alnus kushiroensis Tanai, 6 monocots. Betulaceae and Trochodendroides were not Alnus sp., Fagopsis sp., Dryophyllum sp., Platanus cf. P. recovered. newberriana Heer, Craigia oregonensis (Arn.) Kvaček, Especially in aquatic and helophytic assemblages the Bůžek & Manchester, “Pterospermites” auriculaecordatus Raichikha flora is similar to that of the Paleocene and Early Hollick, Archeampelos acerifolia (Heer) McIver & Basin- Eocene Zaissan Lake basin (South East Kazakhstan) (Fig. 2, ger and others. loc. 8). The Raichikha assemblage includes (Figs 5–7) The flora from Due village (North-Western Sakhalin, Marchantites, Cyclosorus, Osmunda, Woodwardia, Sal- Fig. 3, loc. 12) was first described by Kryshtofovich (1936) vinia, Regnellidium, Taxodium, Nelumbo, Nuphar, Hibis- and Borsuk (1956). The fossils belong to the Conglomerate cus, Limnobiophyllum, Zingiberopsis, Peltandra, Typha, Formation, but mainly the Lower Due Formation (con- Lauraceae and Fabaceae along with Rhamnaceae glomerates, sandstones, silts and coal). The main compo- (Guoania, Ziziphus). The Raichikha flora occupied more nents of this flora are Osmunda sachalinensis Krysht, elevated habitats too, accompanied by ferns, such as Ly- Lygodium asiaticum Borsuk, Ginkgo adiantoides (Ung.) godium and Anemia. The flora includes many evergreen Heer, Metasequoia occidentalis (Newb.) Chaney, Taxo- plants (Magnolia, Ocotea, Neolitsea, Ficus, some Fa- dium dubium (Sternb.) Heer, Glyptostrobus europaeus baceae, Myrtophyllum, Delavaya, etc.) (see Figs 5–7). (Brongn.) Ung., Trochodendroides arctica (Heer) Berry, The Raichikha flora arose in the northern part of the various Amentiflorae (Populus celastrophylla Baikov- subtropical realm of Eastern Eurasia. In the Early Eocene skaya & Sycheva, Myrica ovalifolia Borsuk, Castanea the Raichikha flora suggests that the northern boundary of longifolia Borsuk, Quercus spp., Alnus onorica Borsuk

Figure 5–7. Plant macrofossils showing the distinctive nature of the Early Eocene subtropical flora from Raichikha (Zeya-Bureya sedimentary basin, Amur River Area) (coll. No. 3816, Geological Institute RAS). Figure 5. A, B, F, K – Anemia elongata (Newberry) Knowlton: A – No. 3816/36 (× 2), 2 – No. 3816/128 (× 2), F – No. 3816/426 (× 2), K – No. 3816/371 (×2).•C–Filices sp., No. 3816/105 (× 2.5). • D, E – Taxodium dubium (Sternb.) Heer: D – No. 3816/59 (× 1), E – No. 3816/165 (× 2).•G–Guoania grubovii Fedotov, No. 3816/1996 (× 1). • H, L – Salvinia praeaculeata Berry: H – No. 3816/256 (× 5), L – No. 3816/27(× 5). • I, M – Ziziphus raitschichensis Fedotov: I – No. 3816/469 (× 1), M – No. 3816/113 (× 1). J – Ocotea beringiana Budants., No. 3816/345 (× 1).•N–Morus sp., No. 3816/150 (× 1).

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87 Bulletin of Geosciences • Vol. 85, 1, 2009 and others), Magnolia kryshtofovichii Borsuk, Platanus Hamamelidaceae (Hamamelis, Fothergilla, Corylopsis), spp., Liquidambar europaea A. Braun, Vitis sachalinensis Lauraceae (Sassafras, Daphnogene) and Magnoliaceae Krysht. and others (Sorbaria, Craigia, Zizyphus, Rhamnus, (Akhmetiev 1993). Viburnum, Rosaceae, Daphne, Vaccinium ). This assem- blage consists of more then 40 species. Two other floras (Western Sakhalin, Lower and Middle Late Lutetian–Bartonian warm temperate to Snezhinka Subformations, Fig. 3, loc. 13 and South-East- temperate floras of the North-Western Pacific ern Sakhalin, Naibuti Formation, Fig. 3, loc. 14) were stud- coast (Fig. 4, localities 16, 17), Sakhalin (Fig. 4, ied by Sycheva (1977), Akhmetiev (1993) and Kodrul locality 18), South Primorie (Fig. 4, locality 20) (1999). Both floras are the same age, probably Late Ypresian–Lutetian, but they are distinguished by ecologi- On the North-Western Pacific coast there are several loca- cal factors. The ferns are few (Osmunda sachalinensis lities. Shestakovo (Fig. 4, loc. 16) on the south coast of Krysht., Onoclea hesperia R.W. Brown), gymnosperms Penzhina bay is situated between the mouth of the Shesta- are dominant, especially Ginkgo adiantoides (Ung.) Heer, kovo River and Lovaty Creek. Plant fossiliferous strata Glyptostrobus europaeus (Brongn.) Heer, Metasequoia contain interbedded silts and siltstones, coals and coaly occidentalis (Newb.) Chaney, Taxodium dubium (Sternb.) silts, rare sandstones (300 m), thin coal and coaly clays len- Heer, more rarely Pseudolarix, Sciadopitys, Pinaceae and ses and diatomitic clays with marine diatoms, probably Cupressaceae are almost absent. The main components of Middle Eocene. Coaly clays and silts include fossils (lea- angiosperms are Trochodendraceae, Platanaceae, Faga- ves, seeds, fruits, sometimes conifer woods). ceae, Betulaceae, Salicaceae, Ulmaceae: (Trochoden- This flora was studied by Kryshtofovich (1958), droides arctica (Heer) Berry, Nordenskioldia borealis Ablaev (1985), and Akhmetiev & Bratsteva (1989). The Heer, Platanus acutiloba Borsuk, P. mabuti Oishi & Hu- authors collections include more than 20 species – typical zioka, P. zhuravlevii Medjul., Castanea longifolia Borsuk, elements of the Middle Eocene North-East Asia floras: C. sachalinensis Kodrul, Fagopsis nipponica Tanai, Alnus Equisetum arcticum Heer, Dennstaedtia americana ezoensis Tanai, A. hokkaidoensis Tanai, A. kushiroensis Knowlt., Osmunda macrophylla Penh., Ginkgo ex gr. Tanai, Carpinus kushiroensis Tanai, Populus budantsevi adiantoides (Ung.) Heer, Picea sp. (seeds), Pinus sp. Ablaev, P. celastrophylla (Baikovskaya) Sycheva, P. (leaves), Pinus sp. (seeds), Pseudolarix sp. (bractea), gigantophylla Budants., Ulmus ezoana (Oishi & Hu- Taxodium tinajorum Heer, Metasequoia occidentalis zioka) Kodrul, U. harutoriensis Oishi & Huzioka, (Newb.) Chaney, Sequoia sp., Glyptostrobus europaeus Zelkova kushiroensis Oishi & Huzioka). Tiliaceae are (Brongn.) Heer, Cupressaceae gen., Magnolia praekobus represented by Craigia oregonensis (Arnold) Kvaček, Budants., Trochodendroides arctica (Heer) Berry, Nyssi- Bůžek & Manchester, Plafkeria basiobliqua (Oishi & dium arcticum (Heer) Iljinskaya, Alnus protophylloides Huzioka) Tanai, various Rosaceae, Aceraceae, Ana- Budants. & Golovneva, Myrica beringiana Budants., cardiaceae, Vitaceae (Crataegus sp., Rubus fretalis Juglans pogozheviii Chelebaeva, Craigia oregonensis (Ar- Borsuk, Acer cf. kushiroanum Tanai, Acer cf. palaeo- nold) Kvaček, Bůžek & Manchester, Acer sp., Aesculus platanoides Endo, Acer spp. (fruits), Cotinus sp., Rhus magnifica (Knowlt.) Iljinskaya, Archeampelos acerifolia angustifolia Kodrul, Vitis sachalinensis Krysht., Vitis (Newb.) McIver & Basinger [= “Acer” arcticum (Heer) spp.) and others (more than 100 species). More warmer R.W. Brown], Cissites ochotensis Krysht. elements of these floras are Byttneria iizimae Tanai, The Irgirnin assemblage is very similar to that from Kleinhovia basitruncata (Oishi & Huzioka) Tanai, Ficus Shestakovo. L.Y. Budantsev described the flora from Pod- spiculifolia Nagai, Phytocrene sp. A more ancient ele- kagernaya Bay on the coast (Fig. 4, loc. 17), but it is proba- ment of these floras is Archeampelos acerifolia (Heer) bly younger, late Middle Eocene–(?)Late Eocene. It in- McIver & Basinger. cludes Cercidiphyllum intermedium Budats., Ulmus The flora from the lower part of the coal-bearing compacta Fotjanova, Liquidambar brevilobata Budants., Uglovaya Formation (Artjem coal-field) (Fig. 3, loc. 15) Fagus napanensis Fotjanova, Castanea kamtschatica Fot- consists of typical Early and Middle Eocene components janova, Juglans protocinerea Iljinskaya & Budants. Most (Ginkgo, Taxodiaceae, Trochodendroides, Platanaceae, of the components are the same as Shestakovo (Equisetum Ulmaceae, ancient Amentiferae). There are more often arcticum, Dennstaedtia americana, Ginkgo ex gr. adian-

Figure 6. A, J–L – Ocotea beringiana Budants.: A – No. 3816/571 (× 1.2), J – 3816/179 (× 1), K – No. 3816/387 (× 1), L – No. 3816/406 (× 1). •B–?Tilia sp., No. 3816/486 (× 1). • C – Lindera rajczichensis Fedotov, No. 3816/215 (× 1). • D – Neolitsea rajczichensis Fedotov, No. 3816/443 (× 1). • E, H – Urticaceae gen. et sp. indet.: E – No. 3816/192 (× 1), H – No. 3816/201 (× 1). • F, I – Hibiscus sp.: F – No. 3816/276 (× 1.5), I – No. 3816/209 (× 1). • G – Zingiberopsis magnifolia (Knowlton) Hickey, No. 3816/101 (× 1).

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89 Bulletin of Geosciences • Vol. 85, 1, 2009 toides (Ung.) Heer, Metasequoia occidentalis, Glyp- gen., Hamamelis kushiroensis Tanai, Vitis sp., Tilia todtrobus europaeus, Trochodendroides ex gr. arctica, eojaponica Endo, Mallotus hokkaidoensis Tanai, Rhus Alnus protophylloides (Budantsev 1997). sp., Fraxinus sp. The floristic assemblage is characterized The late Middle Eocene floristic section of Middle by small leaves (up to 3–5 cm long) linked to a winter wet Sakhalin, situated near Uglergorsk town (Snezhinka River climate. basin) (Fig. 4, loc. 18) is very important. The late Middle Eocene Upper Snezhinka subformation represents coals, coal bearing silt and siltstones and rare sandstones (300 m). Paleocene and Eocene floras The fossil leaves and fruits are smaller in size than in the of the Sikhote-Alin volcanic belt (Fig. 1, Kamchatka-Koryakia region, but the taxonomic composi- localities 7–10, Fig. 4, localities 19, 21) tion is similar (Akhmetiev 1993, Kodrul 1999). T.M. Kodrul described 36 species, mostly warm-temperate ele- The Sikhote-Alin volcanic belt is limited to the east by the ments. There are 6 ferns: Osmunda, Dennstaedtia, Tatarian strait and the coast of Japan. The belt experien- Dryopteris, Lastrea, Onoclea, Woodwardia. Conifers (not ced two stages of volcanic activity: Senonian-Paleocene more then 10% specimens) are represented by Pseudo- and late Middle Eocene-Early Miocene. These volcanic larix, Glyptostrobus, Metasequoia, Taxodium. Angio- sequences are separated by gaps. In the southern part of sperms (more than 20 species) include Trochodendroides the volcanic belt sedimentary lenses with floras accumu- arctica (Heer) Berry, Nyssidium arcticum (Heer) Iljin- lated in intermontane valleys and small lake depressions skaya, Platanus sp., various Amentiferae (Alnus ezoensis above the sea-level (Takhobe and Zerkalnaya localities, Tanai, A. ishikariensis Tanai, Alnus spp. (leaves and cat- Fig. 1, locs 9, 10). In the northern part (Malo-Mikhailovka kins), Betula sp., Corylus palaeomaximovicziana (Endo) locality, Fig. 1, locs 7, 8) fossiliferous layers accumulated Tanai, Myrica vindobonensis (Ett.) Heer, Populus in swampy depressions near volcanoes. Their Danian celastrophylla (Baikovskaya) Sycheva, P. gigantophylla or Selandian age is based on radiometric data and compa- Budants, P. grandifolia Endo), some Ulmaceae and Rosa- risons of these floras with the Tsagayan floras in the ceae, Archeampelos acerifolia (Heer) McIver & Basinger, Zeya-Bureya depression. Most typical common elements Acer sp., Euonymus denticulata Kodrul and others. Some are Ginkgo, Taxodium, Cupressinocladus, Trochoden- of the species was described by Tanai (1967, 1970) from droides, Nyssidium, Nyssa, Tiliaephyllum, platanoids etc. Hokkaido. Elements which differ from the typical Tsagayan flora are The Middle Eocene floristic layers were studied on diverse conifers, various Amentiferae e.g., Alnus and Be- Rechnoi peninsula (north of Vladivostok) (Fig. 4, loc. tula (fruits and leaves) (Takhobe and Zerkalnaya locali- 20). The terrigenous Nadezhdinskaya Formation (pre- ties), Corylites and Palaeocarpinus (Malo-Mikhailovka dominantly silts and siltstones with rare sandstones) is locality), Juglandaceae and Fagopsis (Takhobe). Ulmus subdivided into three members (Akhmetiev 1993). The furcinervis is a common component of the Kivda flora Lower Member contains rare fossil plants [Dryopteris sp., (Bureya River Area, Fig. 1, loc. 5 same locality as Tsa- Metasequoia occidentalis (Newb.)] Chaney, Ulmus sp., gayan flora) and the main dominant in the Takhobe and Populus iljinskajaa Akhmetiev, Platanus sp., Lauraceae Zerlalnaya floras. Sorbus and Davidia also occur there. gen. The Middle Member has a richer floristic assem- These Paleocene assemblages typically have leaves blage. It includes Metasequoia and Trochodendroides and fruits of large size. Lignites and coaly clays at the (dominants), Magnolia sp., Alnus cf. protohirsuta Endo, Malo-Mikhailovka locality accumulated in swampy Ostrya sp., Juglandaceae gen., Ulmus ex gr. longifolia depressions and small river systems and yielded Muscites, Ung., Zelkova kushiroensis Oishi & Huzioka, Hamamelis Equisetum, Onoclea, Dennstaedtia, Asplenium, Ginkgo, sp., Leguminosae gen., Cinnamomum sp., Cedrela Fokieniopsis, Cryptomerites, Amurocyparis, Palaeocar- sp., Plafkeria sp. The Upper Member is the main floristic pinus, Corylites, Trochodendroides, Trochodendrocar- horizon. The dominant elements are Metasequoia pus. occidentalis (Newb.) Chaney, Populus iljinskajae The oldest Eocene floras of the Eastern Sikhote-Alin Akhmet., Zelkova zelkovaefolia (Ung.) Bůžek & Kotlaba, basaltic belt are found on the north coast at Sonye Bay Platanus aceroides Goepp., Laurophyllum sp. The rarer (Fig. 4, loc. 19), 200 km south of Sovietskaya Gavan town. elements are Trochodendroides arctica (Heer) Berry, Thin ash lenses between basaltic flows (Kizi Group) con- Nordenskioldia borealis Heer, Liquidambar miosinica tain different gymnosperms (more than 50% of specimens) Hu & Chaney, Cinnamomum sp. and Plafkeria sp. The [Ginkgo, Pinus, Picea, Metasequoia, Taxodium, Glypto- rarest elements are Osmunda sakhalinensis Krysht., strobus, Cryptomeria, Metasequoia, Taxodium, Cunning- Ginkgo adiantoides (Ung.) Heer, Pinus sp., Taxodium hamia, Torreya, (?)Cupressaceae, (?)Podocarpus and dubium (Sternb.) Heer, Zingiberaceae gen., Magnolia sp., others]. Angiosperms are represented by Quercus ex gr. Carpinus ex sect. Eocarpinus, Celtis sp., Leguminosae Dentatae, Fagus sp., Acer ezoanum Oishi & Huzioka and

90 Mikhail A. Akhmetiev • Paleocene and Eocene floristic and climatic change in Russia and Northern Kazakhstan

A BDC E

F G H

I JK

Figure 7. A–Rhus sp., No. 3816/70 (× 2).•B–Ficus schimperi Lesq., No. 3816/417 (× 1).•C–Delavaya fraxinifolia Fedotov, No. 3816/287 (× 2). •D–Leguminocarpon sp. (associated with leaves of Stenolobium), No. 3816/389 (× 3). • E, H, I, J – Celtis sp. E – No. 3816/139 (× 1), H – No. 3816/140 (× 1), I – No. 3816/178 (× 1), J – No. 3816/144 (× 1).•F–Lindera raiczichensis Fedotov, No. 3816/372 (× 1).•G–Stenolobium sp., No. 3816/6 (× 1). • K – Leguminosites raiczichensis Fedotov, No. 3816/251a (× 3).

91 Bulletin of Geosciences • Vol. 85, 1, 2009 others. The abundance of conifers is connected with an up- Acknowledgments land habitat of the assemblage (Akhmetiev 1993). A very similar flora was described by Akhmetiev The author thanks M. Collinson and Z. Kvaček for useful advice (1993) from near the Svetlovodnaya River valley (Fig. 4, and help. The study was supported by Russian Foundation for loc. 21). This flora contains Ginkgo, Metasequoia, Trocho- Basic Research, project N 08-05-00548, SS-4185.2008.5 and dendroides, Fagopsis, Ulmaceae gen., Craigia, Plafkeria, Programme N15 Presidium of the Russian Academy of Sci- Vitis, Acer and others. ences.

Conclusions References

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